The World Flora Online

Herbs, subshrubs, or shrubs. Stems mostly fleshy. Leaves alternate, opposite, or verticillate, usually simple; stipules absent; leaf blade entire or slightly incised, rarely lobed or imparipinnate. Inflorescences terminal or axillary, cymose, corymbiform, spiculate, racemose, paniculate, or sometimes reduced to a solitary flower. Flowers usually bisexual, sometimes unisexual in Rhodiola (when plants dioecious or rarely gynodioecious), actinomorphic, (3 or)4-6(-30)-merous. Sepals almost free or basally connate, persistent. Petals free or connate. Stamens as many as petals in 1 series or 2 × as many in 2 series. Nectar scales at or near base of carpels. Follicles sometimes fewer than sepals, free or basally connate, erect or spreading, membranous or leathery, 1- to many seeded. Seeds small; endosperm scanty or not developed.

Seeds mostly minute, usually with fleshy endosperm; embryo straight Fruit follicular, membranous or leathery, often surrounded by the persistent membranous corolla, opening on the adaxial side Stamens as many or twice as many as the petals, if few then alternate with the petals, slightly perigynous; filaments free; anthers 2-celled, introrse, dehiscing longitudinally Petals the same number as the sepals, free or variously connate, hypogynous Carpels superior, the same number as the petals free or united at the base, 1-celled; ovules many or rarely few, inserted on the adaxial suture; styles short or elongated Hypogynous scales present within the stamens Flowers actinomorphic, hermaphrodite, usually cymose Sepals free or united into a tube, often 4 or 5 Herbs and undershrubs, usually succulent; leaves opposite or alternate, exstipulate Fruit follicular, membranous or leathery, often surrounded by the persistent membranous corolla, opening on the adaxial side Stamens as many or twice as many as the petals, if few then alternate with the petals, slightly perigynous; filaments free; anthers 2-celled, introrse, dehiscing longitudinally Petals the same number as the sepals, free or variously connate, hypogynous Carpels superior, the same number as the petals free or united at the base, 1-celled; ovules many or rarely few, inserted on the adaxial suture; styles short or elongated Hypogynous scales present within the stamens Flowers actinomorphic, hermaphrodite, usually cymose Sepals free or united into a tube, often 4 or 5 Herbs and undershrubs, usually succulent; leaves opposite or alternate, exstipulate

Widely distributed in temperate and tropical regions, especially in mountainous districts, rare in South America and Australasia

Plantes'ordinairement charnues, le plus souvent herbacées, parfois un peu ligneuses, rarement arbustives.'Feuilles'opposées, verticillées ou alternes, généralement succulentes, simples, entières ou faiblement découpées, rarement lobées ou imparipennées, dépourvues de stipules.'Inflorescences'cymeuses, axillaires ou terminales.'Fleurs'☿, rarement ♂♀, régulières, hétérochlamydées, 5(3-32)-mères; sépales libres ou soudés, souvent un peu inégaux; pétales libres ou soudés en tube; androcée libre ou soudé au tube de la corolle, obdiplostémone ou haplostémone par avortement du verticille épipétale; anthères dithèques introrses; gynécée à carpelles en nombre égal à celui des enveloppes florales, libres ou un peu soudés à la base, généralement munis chacun à la base d'un appendice squamuleux (nectarifère ? de forme variable et de petites dimensions; styles subulés, à stigmate capité; ovules anatropes, petits, nombreux, rarement ( Crassula ) 1-2 par carpelle, insérés sur la suture ventrale.'Fruits'folliculaires, membraneux ou coriaces, s'ouvrant par la suture ventrale.'Graines le plus souvent petites et légères, à tégument lisse, ridé ou finement tuberculé; albumen peu abondant ou nul.\n\t\t\tLes Crassulacées, qui comptent 33 genres et environ 1500 espèces, paraissent le mieux représentées dans les régions subtropicales à longue saison sèche. Elles sont rares sous les tropiques et s'y rencontrent généralement dans les régions d'altitude. Ce sont des xérophytes qui atteignent leur plus grand développement dans les régions à précipitations pauvres (Mexique, Cap, bassin méditerranéen). Plantes à reproduction végétative très aisée, même par bulbilles apparaissant sur les feuilles.

Seeds minute; endosperm usually present; embryo straight Carpels superior, equal in number to the petals, free or slightly connate at the base, 1-locular; ovules (few–)many, inserted on the adaxial suture; style short or elongated, stigmatose on the inner side of the apex Fruit follicular, membranous or leathery, often surrounded by the persistent membranous corolla, opening on the adaxial side Petals the same number as the sepals, free or variously connate Stamens hypogynous or epipetalous, as many as the petals or twice as many, and in 2 whorls, frequently obdiplostemonous; filaments free or adnate to the petals; anthers 2-thecous, introrse, opening by a longitudinal slit; scale-like nectaries usually present between the stamens and carpels Flowers usually in terminal cymose inflorescences, less often in spikes or racemes or solitary in the leaf-axils, with or without bracts, regular, hermaphrodite, rarely unisexual and dioecious, mostly (3–)4–5(–±30)-merous Sepals free or united into a tube, persistent Annual, biennial or perennial herbs, rarely small shrubs, usually succulent Leaves opposite or alternate, rarely verticillate, exstipulate Carpels superior, equal in number to the petals, free or slightly connate at the base, 1-locular; ovules (few–)many, inserted on the adaxial suture; style short or elongated, stigmatose on the inner side of the apex Fruit follicular, membranous or leathery, often surrounded by the persistent membranous corolla, opening on the adaxial side Petals the same number as the sepals, free or variously connate Stamens hypogynous or epipetalous, as many as the petals or twice as many, and in 2 whorls, frequently obdiplostemonous; filaments free or adnate to the petals; anthers 2-thecous, introrse, opening by a longitudinal slit; scale-like nectaries usually present between the stamens and carpels Flowers usually in terminal cymose inflorescences, less often in spikes or racemes or solitary in the leaf-axils, with or without bracts, regular, hermaphrodite, rarely unisexual and dioecious, mostly (3–)4–5(–±30)-merous Sepals free or united into a tube, persistent Annual, biennial or perennial herbs, rarely small shrubs, usually succulent Leaves opposite or alternate, rarely verticillate, exstipulate

Fls regular, hypogynous or nearly so, (3)4–5-merous or occasionally more, usually perfect; sep, pet, and pistils normally of the same number, the stamens as many or twice as many; sep distinct or ± united at base; pet in our genera distinct or very shortly connate below; carpels generally distinct or nearly so and ripening into follicles; a small nectariferous appendage borne externally on each carpel near the base; succulent herbs or shrubs with fleshy, simple lvs and usually cymose fls. 25/900.

SELECTED REFERENCES Berger, A. 1930. Crassulaceae. In: H. G. A. Engler et al., eds. 1924+. Die natürlichen Pflanzenfamilien..., ed. 2. 26+ vols. Leipzig. Vol. 18, pp. 352-485. Britton, N. L. and J. N. Rose. 1903. New or Noteworthy North American Crassulaceae. New York. [Preprinted from Bull. New York Bot. Gard. 3: 1-45. 1903.] Britton, N. L. and J. N. Rose. 1905. Crassulaceae. In: N. L. Britton et al., eds. 1905+. North American Flora.... 47+ vols. New York. Vol. 22, pp. 7-74. Eggli, U., ed. 2003. Illustrated Handbook of Succulent Plants. Crassulaceae. New York. Ham, R. C. H. J. van and H. ’t Hart. 1994. Phylogenetic relationships in the Crassulaceae inferred from chloroplast DNA restriction-site variation. In: R. C. H. J. van Ham. 1994. Phylogenetic Implications of Chloroplast DNA Variation in the Crassulaceae. Utrecht. Pp. 81-103. Ham, R. C. H. J. van and H. ’t Hart. 1998. Phylogenetic relationships in the Crassulaceae inferred from chloroplast DNA restriction-site variation. Amer. J. Bot. 85: 123-134. ’t Hart, H. 1995. Infrafamilial and generic classification of the Crassulaceae. In: H. ’t Hart and U. Eggli, eds. 1995. Evolution and Systematics of the Crassulaceae. Leiden. Pp. 159-172. Knapp, U. 1994. Skulptur der Samenschale and Gliederung der Crassulaceae. Bot. Jahrb. Syst. 116: 157-187. Mauritzon, J. 1933. Studien über die Embryologie der Familien Crassulaceae und Saxifragaceae. Lund. Mayuzumi, S. and H. Ohba. 2004. The phylogenetic position of eastern Asian Sedoideae (Crassulaceae) inferred from chloroplast and nuclear DNA sequences. Syst. Bot. 29: 587-598. Mort, M. E. et al. 2001. Phylogenetic relationships and evolution of the Crassulaceae inferred from matK sequence data. Amer. J. Bot. 88: 76-91. Ohba, H. 1978. Generic and infrageneric classification of the Old World Crassulaceae. J. Fac. Sci., Univ. Tokyo, Sect. 3, Bot. 12: 139-198. Quimby, M. W. 1939. The Floral Morphology of the Crassulaceae. Ph.D. dissertation. Cornell University. Spongberg, S. A. 1978. The genera of Crassulaceae in the southeastern United States. J. Arnold Arbor. 59: 198-248. Uhl, C. H. 1963. Chromosomes and phylogeny of the Crassulaceae. Cact. Succ. J. (Los Angeles) 35: 80-84.

Annual, biennial, or perennial herbs or shrubs [rarely scandent], succulent. Stems mostly alike and all potentially flowering, (flowering stems annual, axillary, overtopping primary stem). Leaves usually persistent, sometimes deciduous or caducous, cauline or sometimes in dense basal rosettes, alternate, opposite, or whorled, simple (pinnate in Bryophyllum), (heteromorphic in Dudleya, Echeveria, Graptopetalum); stipules absent; petiole usually absent; blade margins usually entire, sometimes toothed or lobed. Inflorescences terminal or axillary, spikes, thyrses, or mostly cymes, branches commonly cincinni (scorpioid cymes) and at first commonly coiled, or rarely flowers solitary. Flowers bisexual (sometimes unisexual in Rhodiola), radially symmetric (except sometimes calyx); perianth and androecium hypogynous or weakly perigynous; hypanthium present; sepals (3-)4-5(-12)[-30+], distinct or connate basally (or into tube); petals (3-)4-5(-12)[-30+], distinct or connate basally or sometimes into tube, margins entire (fimbrillate in Jovibarba, crenate in Phedimus); nectaries as scales at base of carpels; stamens as many as sepals or 2 times as many and in 2 series, antipetalous if in 1 series, free or adnate to corolla base (or tube); anthers dehiscent longitudinally and laterally; pistils (3-)4-5(-12)[-30+] [rarely fewer than sepals], mostly distinct or nearly so, (3-)4-5(-12)[-30+]-carpellate; ovary superior or semi-inferior, 1-locular; placentation submarginal; ovules anatropous, bitegmic, crassinucellate or tenuinucellate; styles (3-)4-5(-12)[-30+], terminal, distinct; stigmas (3-) 4-5(-12)[-30], terminal, capitate. Fruits usually whorls of follicles, each dehiscent along adaxial suture, rarely whorls of flimsy 1-seeded utricles (Sedella) or capsules of basally connate pistils (Diamorpha, Jovibarba). Seeds 1-20+ per carpel, brownish, fusiform or ellipsoid; embryo straight; endosperm oily, scant.

Hypogynous scales (scale-like nectaries) usually present, small, applied to the base of the outer face of the carpels Ovary superior; carpels equal in number to the petals, free or united (up to the middle); ovules many or few to one in each carpel, anatropous, inserted on the adaxial suture; styles short, sometimes nearly absent or ± elongate; stigmas capitate Stamens as many or twice as many as the petals; filaments free or more or less connate with the corolla-tube; anthers 2-celled, introrse, basifixed, dehiscing longitudinally Seeds mostly minute with smooth, rugose or tuberculate tegument and usually fleshy endosperm; embryo straight Fruit follicular, membranous or leathery, opening along the free part of the adaxial suture Petals as many as the sepals, free or ± connate, hypogynous Sepals free or ±: united at the base, sometimes in a ± long tube Flowers (3)4-5(6-32)-merous, actinomorphic, usually bisexual, in axillary or terminal cymes often grouped in corymb- or panicle-like inflorescences, less often in racemes or spikes or solitary in the leaf-axil Leaves opposite, verticillate or alternate, usually simple, sometimes compound, exstipulate, often ± thick and fleshy-succulent Ovary superior; carpels equal in number to the petals, free or united (up to the middle); ovules many or few to one in each carpel, anatropous, inserted on the adaxial suture; styles short, sometimes nearly absent or ± elongate; stigmas capitate Stamens as many or twice as many as the petals; filaments free or more or less connate with the corolla-tube; anthers 2-celled, introrse, basifixed, dehiscing longitudinally Seeds mostly minute with smooth, rugose or tuberculate tegument and usually fleshy endosperm; embryo straight Fruit follicular, membranous or leathery, opening along the free part of the adaxial suture Petals as many as the sepals, free or ± connate, hypogynous Sepals free or ±: united at the base, sometimes in a ± long tube Flowers (3)4-5(6-32)-merous, actinomorphic, usually bisexual, in axillary or terminal cymes often grouped in corymb- or panicle-like inflorescences, less often in racemes or spikes or solitary in the leaf-axil Leaves opposite, verticillate or alternate, usually simple, sometimes compound, exstipulate, often ± thick and fleshy-succulent